SPINAL LESIONS Pathology
In this condition, there is degeneration of dorsal nerve roots, central
I. Syringomyelia
Pathology
In this condition, there is excessive overgrowth of neuroglial tissue
(gliosis) with cavity formation. It involves the grey matter around
the central canal. There is dilatation of the central canal of the spinal
cord.
Cervical enlargement is the commonest site; therefore, hands and
arms most affected.
In other words, in this condition, there is destruction of the cord
involving central canal and its surrounding area due to formation
of elongated cavities.
Features
Since the disease starts around the central canal, the lesion involves
the decussating spinothalamic fibres in the anterior white
commissure. Therefore, there is bilaterlal loss of pain and
temperature sensations below the lesion but other sensations are
preserved in the uncrossed tracts of the posterior columns. Thus the
condition results in what is called as dissociated sensory loss
(dissociated anaesthesia)
The defect is bilateral and usually occurs in the hands and arms due
to the predilection of syringomyelia for the cervical enlargement of
the cord.
#The gliosis and cavitation may spread to involve the anterior horn
cells, causing flaccid paralysis of the upper limb muscles. As the
lesion spreads to the adjacent corticospinal tract, it results in upper
motor neuron paralysis of the legs.
As the disease advances, various other tracts and areas are
destroyed. For example, Romberg’s sign (see below) can eventually
appear pointing that posterior column has been involved.
II. Tabes dorsalis
to the dorsal root ganglion. It is one of the manifestations of neuro-
syphilis ( a type of tertiary syphilis).
Features
1. Pain: There is ‘lightening’ pain; this is the first effect of the
disease. It is due to the stimulation of the pain fibres in the dorsal
root.
2. Later, pain is lost. Due to this, there can be joint deformity.
This is because mild injury is neglected (due to absence of pain);
therefore, repeated injury damages the articular surfaces resulting in
deformed joints (Charcot’s joints).
3. There is loss of dorsal column sensations:
i) There is loss of position sense, vibratory sense, sense of
stereognosis and discriminative touch
ii) The loss of proprioception (or position sense0 results in
sensory ataxia.
4. Romberg’s sign
This is used to differentiate between cerebellar ataxia and sensory
ataxia.
The person is asked to close his eyes and stand straight with his feet
close together. A normal person can keep his balance with the eyes
closed, using the proprioceptors in the legs
Sensory ataxia:
A person who has lost proprioception in the legs does not know
where the ground is relative to the legs. He can maintain his balance
using his eyes; however, on closing his eyes, he is unable to balance
himself and starts swaying
Cerebellar ataxia:
A person suffering from cerebellar ataxia is unable to stand erect
(with his feet together) even when his eyes are open. In other words,
he sways irrespective of whether the eyes are open or closed.
 Note: if there is infection, malnutrition etc., then the duration gets
Note: Romberg’s sign is positive in sensory ataxia; the patient is prolonged.
able to stand erect with his eyes open but not with his eyes closed. Features
III. Transverse section (transection) of the spinal cord Termination Function
(Note: Students may kindly study this topic after we have discussed
End on motor neurons in the anterior horn For spinal reflexes
the chapter on ‘Reflexes’. They may, however, study the sensory
of the spinal cord (directly or through
aspects of the Brown-Sequard syndrome, which was discussed in
interneurons)
our lecture on 24.11.2018)
Ascend up in specific tracts (specific For conscious perception of specific
1. Complete /Incomplete
sensory pathways) to a specific site on the sensations
2. Hemisection (Brown-Sequard syndrome)
sensory cortex
1. Complete/incomplete
The severity of the problem depends upon the site of the lesion. Ascend up in non-specific sensory For alert state
Higher up the lesion in the spinal cord, greater is the extent of pathways (Ascending reticular activating
damage. system or ARAS) and go to the entire
i) For example, transection of the cervical cord just below C5 neocortex
results in quadriplegia Ascend up to cerebellum For unconscious proprioception
ii) Transection between cervical and thoraco-lumbar vertebra Ascend up to tectum (of the midbrain) For visuo-spinal reflexes
results in paraplegia
During the stage of spinal shock:
iii) Transection above C5 is fatal because of paralysis of respiratory
i) there is flaccid (LMN type) paralysis both at the level as well as
muscles (recall that phrenic nerve (C3,4 and 5) supplies the diaphragm)
below the level of lesion
stages
ii) muscle tone is decreased
1. Stage of spinal shock
iii) reflexes are decreased
2. Stage of reflex hyper-activity
iv) no sensation
Stage of spinal shock
Stage of reflex hyperactivity
After spinal cord section, there is a time period wherein the peripheral
Features
nerve fibres do not conduct impulses. Thus, the nerve fibres, although
i) reflexes are increased
anatomically present, are not functional. This stage is known as the stage
ii) muscle tone is increased
of spinal shock. The cause of this is not exactly known.
iii) there is spastic (‘UMN’ type) paralysis below the level of the
Duration of spinal shock:
lesion (at the level of the lesion, flaccid paralysis persists)
The duration of spinal shock depends upon the degree of encephalization:
iv) Babinski’s sign
i) frogs : 2 to 3 minutes
v) Clonus
ii) dogs/cats : a few hours
Note: If a complete transection results in paraplegia, the lower limbs are
iii) human beings : 3 weeks or more
held in a slightly flexed position; this is therefore known as ‘paraplegia
flexion’. This is because the flexors are more hypertonic than Column Ascending tracts Descending tracts
the extensors. Anterior white column Anterior spinothalamic Anterior corticospinal,
If an incomplete transection results in paraplegia, there is vestibulospinal, tectospinal,
‘paraplegia in extension’. This is because in this medial reticulospinal
condition, there is more extensor hypertonia.
Lateral white column Lateral spinothalamic, anterior Lateral corticospinal,
2. Brown-Sequard syndrome
This is the name given to the condition resulting from a
spinocerebellar, posterior rubrospinal, lateral
hemi-section of the spinal cord spinocerebellar, spinotectal reticulospinal, spinal
Features sympathetic
1. Pain and temperature are lost on the opposite side of Posterior white column Fasciculus gracilis (tract of -
the lesion Goll) and fasciculus cuneatus
2. All other sensations are lost on the same side of the (tract of Burdach)
lesion ii) Ventral (or anterior) column (VC)
3. Paralysis occurs on the same side of the lesion iii) Lateral column (LC)
4. At the level of the lesion, there is ‘LMN’ type of paralysis iv)
5. Below the level of the lesion, there is ‘UMN’ type of paralysis.
6. Touch is least affected, as it has a dual pathway.
TOPIC: ASCENDING PATHWAYS
Recall that in the spinal cord, dorsal (or the posterior) root is sensory
and the ventral (or the anterior) root is motor. Thus, the sensory
ascending pathways enter the spinal cord through the dorsal root;
thereafter, they ascend in one of the three columns viz. dorsal (or
posterior), lateral and ventral (or anterior).
After entering the spinal cord, the sensory fibres can take the
following routes:
Different tracts in the various columns

In our set of lectures on CNS I, we shall be discussing only the

specific sensory pathways. Based on the above fact, the ascending pathways can be broadly
The specific sensory pathways divided into two main pathways:
As the name suggests, they carry specific sensations. As we already
know, they can ascend in one of the following columns viz. 7. The antero-lateral (or the ventro-lateral) system
i) Dorsal (or posterior) column (DC) The antero-lateral system is further sub-divided into:
 i) Anterior and ii) Lateral
8. The dorsal (or the posterior) column system

System Carries
Anterior Crude touch and crude pressure
Lateral Pain and temperature
Dorsal column Fine touch, tactile localisation, vibration,
proprioception, 2-point discrimination, pressure
discrimination, stereognosis
The systems carry the specific sensations as under:

2-point discrimination: means the ability to recognize (with the eyes

closed) two closely-placed points as separate. It is best in the finger
tips and worst in the back.
Stereognosis : means the ability (with the eyes closed) to recognize
a familiar object placed in the hand just by feeling it e.g. recognizing
a pen, key etc.
Mnemonic
“ PCT does NOT carry PCT!”
The Posterior Column Tract does NOT carry Pain,
Crude touch and Temperature.)
Let us now do in detail, the route taken by the above systems:
3. The antero-lateral (or the ventro-lateral) system
Use the following diagram as the reference for studying the text that
follows immediately: Carefully study the diagram above and note
the following points:
There are 3 separate sensory neurons viz. I, II and III, carrying the
sensation from the receptor to the cortex → they are known
I. First order sensory neurons:
As we already know, the first order sensory neurons enter the spinal
cord through the dorsal root, at their relevant spinal segments. Their
course is from the receptor to the spinal cord. These are pseudo-
unipolar neurons; their cell bodies are located in the dorsal root
ganglion (DRG). Obviously therefore, there is no synapse at the
DRG
II. Second order sensory neurons:
i) The first order sensory neurons synapse with the second
order sensory neurons in the spinal cord;
ii) in the spinal cord, they cross to the opposite side.
iii) The second order neurons (after having crossed to the
opposite side in the spinal cord) ascend up and synapse with certain
nuclei in the thalamus → therefore, the antero-lateral system is also
known as the anterior spino-thalamic tract and the lateral spino-
thalamic tract):
Note:
The first order sensory neurons may ascend up (or descend down) III. Third order sensory neurons
on the same side a few segments in the spinal cord before making a From the thalamus, the third order neurons arise; they finally end in
synapse in the spinal cord and then crossing to the opposite side e.g. the sensory cortex.
the fibres entering at C5 may synapse at C3 as shown in the Sensory cortex:
diagram below: In the sensory cortex, the third-order sensory neurons
end in the post-central gyrus.
i) The post-central gyrus is also known as the primary somato-
sensory area or SI (this corresponds to Brodmann’s area 3,1,2)
ii) There is one more sensory area known as SII; it lies in the
superior lip of the Sylvian fissure
iii) There is another area (the posterior parietal lobule) just
posterior to the post-central gyrus; it is known as the sensory
association area; it corresponds to the Brodmann’s area 5 and 7

The nuclei of the thalamus with which these second order neurons
synapse are known as the ventro-posterior nucleus of the thalamus:
the ventro-posterior nucleus of the thalamus has two parts:

a) ventro-posterior medial nucleus of the thalamus:

This receives sensations from the face (via the trigeminal nerve)
b) ventro-posterior lateral nucleus of the thalamus:
This receives sensations from the rest of the body
(Note:)
Fibres conveying the anterolateral column sensations from all over
the body except the face and head (i.e. spinothalamic fibres) are
called spinal lemniscus.
Fibres conveying the anterolateral column sensations from all the Somatotopy and sensory homunculus
face and head (i.e. spinothalamic fibres) are called trigeminal Each point in the body is represented by a corresponding point in the
lemniscus (since these sensations are carried by the trigeminal post-central gyrus; this is known as somototopy or point-for-point
nerve) representation. Thus, the entire body is represented in a miniature
manner in the post-central gyrus; this is known as the sensory
homunculus. The body is represented upside down.

The area allotted to each body part in the post-central gyrus depends
upon the density of receptors in the area and the degree of
specialization. Thus, the lips, face and thumb have a large
representation whereas the back, forearm etc. have a much smaller
representation.1
4. The The dorsal (or the posterior) column system iv) first order sensory neurons:
Use the following diagram as the reference for studying the text that As we already know, the first order sensory neurons enter the spinal
follows immediately: Carefully study the diagram above and note cord through the dorsal root, at their relevant spinal segments. Their
the following points: course is from the receptor to the spinal cord.
There are 3 separate sensory neurons viz. I, II and III, carrying the vi) Second order sensory neurons:
sensation from the receptor to the cortex → they are known i) The first order sensory neurons ascend up the spinal cord on
(respectively) as the first order, second order and third order the same side and synapse with the second order sensory neurons in
sensory neurons. the medulla;
In the medulla, there are two nuclei on which the first order sensory
neurons end; these are known as the:
iv) Cuneate nucleus
v) Gracile nucleus
1. The first order sensory neurons of the dorsal column
ascending up the spinal cord and carrying sensations from the upper
part of the body are known as → the tract of Burdach (or the
fasciculus cuneatus); they end on the cuneate nucleus.
 Anterolateral column Dorsal column-medial lemniscus system
Carries: pain, temperature, crude touch Carries: others
First order sensory neurons synapse in the spinal cord First order sensory neurons synapse in the
2. The first order sensory neurons of the medulla
dorsal column ascending up the spinal cord and Crossing over: spinal cord Medulla
carrying sensations from the lower part of the Lamination: Fibres from the lower parts of the body Lamination: Fibres from the lower parts of
body are known as → the tract of Goll (or the are placed laterally (see the diagram below)* the body are placed medially (see the
fasciculus gracilis); they end on the gracile diagram below)*
nucleus. Sensations that are conveyed are not so precise w.r.t. Carries better developed sensory modalites
ii) in the medulla, they cross to the their perception in relation to localization or which are precise in location and
opposite side. discrimination discrimination etc.
These fibres which cross to the opposite side
are known as the internal arcuate fibres (IAF); Conducts relatively primitive senses of crude touch,
iii) The second order neurons (after having pain and temperature.
crossed to the opposite side in the medulla) This is composed of much smaller myelinated fibres The system is composed of large, myelinated
ascend up and synapse with certain nuclei in (averaging 4 micrometers in diameter) that transmit nerve fibres that transmit signals to the brain
the thalamus → these ascending fibres are signals at velocities ranging from a few meters per at velocities of 30 to 110 m/sec
known as the medial lemniscus (or Reil’s second up to 40 m/s
band) → hence, the posterior (or the dorsal) Much smaller degree of spatial orientation of the High degree of spatial orientation
column system is also known as the posterior nerve fibres w.r.t. their origin
column-medial lemniscus system. Poor degree of spatial localization High degree
The nuclei of the thalamus with which these Gradations of intensity is poor High
second order neurons synapse are known as the Ability to transmit rapidly changing or rapidly High
ventro-posterior nucleus of the thalamus: the repetitive signals is poor
ventro-posterior nucleus of the thalamus has Crude type of transmission system Fine type of transmission system
two parts: Phylogenetically older Phylogenetically newer
v) ventro-posterior medial nucleus of the
thalamus:
This receives sensations from the face (via the trigeminal nerve)
vi) ventro-posterior lateral nucleus of the thalamus:
This receives sensations from the rest of the body
vii) Third order sensory neurons
From the thalamus, the third order neurons arise; they finally end in
the sensory cortex.
comparison of the 2 major specific sensory systems:
 It is the psychical adjunct of an imperative protective reflex (its
meaning has been explained in the lecture).

Types of pain
Fast Pain Slow pain
Other names Initial pain, first Delayed pain,
pain second pain
Localisation Well localized Poorly
localized
Character Pin-prick Dull, burning
sensation sensation
Fibres carrying ‘A delta’ ‘C’ fibres
the sensation (Group III) (Group IV)
(S = Sacral; L = Lumbar; T = Thoracic; C = Cervical) Associated Produces Produces
Physiology of Pain features withdrawal nausea,
response vomiting
Characteristics:
sweating.
The sensation of pain has the following characteristics:
1. Protective : Origin Usually cutaneous,
cutaneous muscle, joint or
The sensation of pain has protective value; it helps to
(superficial visceral (deep)
protect us from the damaging stimulus. Further, it
pain)
draws attention to underlying disease. Density of Somatic Visceral : less
2. Pre-potent: receptors structures :
This means it ‘blocks’ any other sensation occurring more density of
simultaneously. pain receptors
3. Unpleasant Sites Fast pain
4. Autonomic nervous system symptoms: receptors are
It may be associated with ANS symptoms like present only in
nausea, vomiting, sweating etc. the skin
Referred pain Superficial pain Deep pain :
Sherrington’s definition of pain is not referred Visceral pain
and deep
 somatic pain is
referred
Muscle pain
When the muscles contract, there is accumulation of pain- producing
metabolite (Lewis’ ‘P’ factor); the Lewis’ ‘P’ factor may be
potassium (or any other metabolite). Now, if the blood supply is
adequate, there is not much accumulation of the ‘P’ factor (it gets
‘washed away’). However, in situations of less blood supply (which
is known as ischaemia), there is pain.
Examples
i) Angina pectoris
When the blood supply to the heart is inadequate,
chest pain can occur on exertion
ii) Intermittent claudication
When the blood supply to the legs is inadequate (as
in Buerger’s disease which is a vaso-occlusive
disease), walking results in more accumulation of the
P factor and the person feels pain  on resting for a
while, the P factor gets removed and the pain
decreases again the person can for some distance.
This situation of walking for some distance, taking
rest in between, is known as intermittent claudication
(claudication means limping
Visceral pain
Features
i) It has all the features of slow pain
ii) Distention is a potent stimulus
iii) It is associated with muscle spasm in the muscles
overlying the viscera (there is reflex contraction of the
abdominal muscles). It occurs especially (but not
necessarily) if the peritoneum is involved. This rigidity
of the abdominal muscles is known as ‘guarding’.
iv) Referred pain can occur (explained below)
v) Radiation of pain can occur (explained below)
Receptors for pain
i) These are known as nociceptors and they are the
modified free nerve endings of the pain nerve fibres
ii) They are slowly- adapting (the concept has already been
explained in the earlier lecture notes)
iii) They are ‘polymodal’ i.e. they can be stimulated by
many sensory modalities e.g. thermal, mechanical,
chemical etc.
iv) Density of the pain receptors:
a) It is maximum in the skin; it is also significant in
the joints, periosteum, tentorium, falx cerbri
b) It is less in the viscera
c) There are no pain receptors in the brain!
d) Receptors for fast pain are present only in the
skin
v) Vanilloid receptors (VR)
These are receptors for pain; they are of many
types:
a) VR-1:
These respond to vanillins (which are
a group of compounds causing pain,
 including capsacin), protons (H+) and
temperature > 43 degree Celsius
b) VRL-1 (VR Like - 1):
These respond to temperatures > 50
degree Celsius (but not to capsaicin)

Note:
The VR belong to a broad family of receptors known as the
transient receptor potential (TRP) channels; these are non-specific
calcium channels. There are several types (and subtypes) of TRP
channels e.g.:
TRPV1: this is same as VR-1 mentioned above
TRPM8: this is same as the cold and menthol receptor (CMR-1); as
the same suggests, it responds to cold and menthol The following table shows the Rexed laminae and their
Pathways corresponding nuclei groups/structures:
Recall that pain (and temperature) are carried by the lateral Lamina Corresponding nuclei/structures
spinothalamic tract (discussed in the earlier lectures). Number
The first synapse for pain is in the spinal cord. I Postero-Marginal Nucleus
Spinal cord II Substantia Gelatinosa Rolando (SGR)
The gray matter of the spinal cord was classified by BrorRexed (a III, IV Nucleus Proprius
neuroscientist) into 10 laminae; these are known as the Rexed
V, VI Base of dorsal column
laminae; the following diagram shows the Rexed laminae:
VII Lateral horn
VIII and IX Groups of nuclei of anterior horn
X Surrounds the central canal (grey commissure
and the substantia gelatinosa centralis)
Lamina VII is further subdivided into:

i) E1 : Intermediate lateral group of the lateral horn

ii) E2 : Intermediate medial group of the lateral horn
iii) E3 : Nucleus dorsalis (Clarke’s column)
iv) Laminae VIII and IX are further subdivided into Important point: Pain can get referred to atypical sites.
i) G1 : Medial group of nuclei of the anterior horn
ii) G2 : Central group of nuclei of the anterior horn When pain occurs in the visceral structure as well as is referred to a
iii) G3 : Lateral group of nuclei of the anterior horn somatic structure, the pain seems to move from the visceral structure
to the somatic structure. This is known as radiation.
Dermatomal rule:
Pain from a visceral structure is referred to a somatic structure
which is innervated by the same spinal segment as the visceral
structure.
Examples:
i) When the gall bladder gets inflamed, it
irritates the diaphragm. The
diaphragm is supplied by the phrenic
nerve (C3,C4,C5); also the tip of the
right shoulder shares the same spinal
segments.
Embryologically, the diaphragm
migrates from the neck region to its
adult location (between the chest and
the abdomen) and therefore, takes its
Referred pain nerve supply with it.
When pain occurs in a somatic structure (at a distance) due to a ii) Embryologically, the heart originates
problem in a visceral structure, it is known as referred pain. from structures in the neck/upper
Examples thorax; therefore, its fibres enter the
i) Gall bladder problem referred as pain at the C3-C5 spinal segments
tip of the right shoulder. iii) Embryologically, the testicle has migrated (along with its
ii) Cardiac pain typically gets referred to the nerve supply) from the urogenital ridge (from which the kidney and
inner aspect of the left arm the ureter have developed)
iii) Appendix pain gets referred to the umbilicus
iv) Ureteric pain gets referred to the testicle Theories of referred pain:
1. Convergence theory

The first order neurons of the somatic and the visceral structure
converge on the same
second order sensory neuron. However, to the brain, it appears to be
coming from the somatic structure.
2. Facilitation theory
3. Collaterals from the visceral afferent cause
subliminal fringe effect on the
4. second order neuron of the somatic afferent.
Gate control theory of pain
The first order sensory neurons synapse in the Rexed lamina II (i.e.
the Substantia Gelatinosa Rolando, SGR) of the spinal cord; from
here, the second order sensory neurons arise. The SGRis a very
important area  in a way, it can be considered as thegate for pain
transmission. If the gate is ‘open’, pain sensation can be transmitted
up; if it is ‘closed”, it will be blocked. As discussed, the
transmission of pain can be modified at this gate.
Transmission at the gate can be modified by:
1. Peripheral input
2. Central input
1. Modification by peripheral inputs:
Stimulation of the touch afferents (from the area
where there is pain) can ‘block’ the transmission of
pain by pre-synaptic inhibition in the SGR.
 Massaging, shaking, acupuncture, TENS
(transcutaneous electrical nerve stimulation) may act
in this fashion
2. Modification by central inputs:
(central inhibition of pain/endogenous pain relief system)
Periaquaductal gray (PAG) of
midbrain

Enkephalin

Raphe magnus nucleus

(via serotinergic neurons)

Serotonin SC = spinal cord; S = serotonin; E = Enkephalin;

Enkephalinergic inter-neurons in spinal cord
Enkephalin
First order neurons for pain in the spinal cord
I and II: first and second order neuron for pain
Endogenous pain relief systems
Endogenous opioids (like beta-endorphins, enkephalins, dynorphins
etc.) relieve pain.
Opioid receptors are present at various sites : PAG, SGR, reticular
formation,thalamus, limbic system etc.
Opioids can act at various sites e.g., at the periphery (at the level of
the receptor), dorsal horn, PAG.
(Stress analgesia: As the name suggests, there is decrease of pain in
situation of stress! For example, fighting injured soldiers at the
battle field do not feel pain. This may be due to release of opioid
peptides.

Pain relief mechanisms

A. Modification at the gate:
 Transmission at the gate (at the SGR in the dorsal - At the site of pain : by blocking the ‘gate’
horn) can be decreased by stimulating the touch - Away from the site of pain : by release of opioids
afferents from the pain producing area; these touch C. Drugs
afferents could decrease the pain transmission at the D. Surgical
gate. - Cutting the nerve
- Posterior rhizotomy
- Anterolateral cordotomy
- Medullary cordotomy
- Thalamotomy
- Frontal lobotomy

RECEPTORS

I. Introduction
A receptor basically functions like a receiver/transducer:
It converts different forms of energies viz. mechanical, chemical
(e.g. smell, taste), electromagnetic (e.g. vision) into electrical energy
i.e. action potential. (The ‘common currency’ in the body is the
action potential; all different forms of stimuli need to be converted
The following procedures may act by modifying the transmission of into action potentials for transmission).
pain at the gate: (Transduction means conversion of one form of energy to another)
- Shaking/massaging the injured area A receptor can either be:
- Counter-irritants i) A specialised cell, lying adjacent to a sensory
- Acupuncture at the site of injury neuron (e.g. hair cells for hearing)
- TENS (transcutaneous electrical nerve ii) nerve ending can itself be modified to act as the
stimulation) receptor; in other words, in such cases, the
‘naked’ or the ‘free’ nerve endings themselves
function as the receptor ( e.g. receptors for pain
B. Acupuncture:
This may act in two ways:
 II. Classification of receptors:
A. Based (essentially) on the site from where the information is
conveyed :

Type Sense Example

Tele-receptors Events at a distance Vision, hearing
Exteroceptors Events close to the body Smell
Interoceptors The internal environment pH, pO2
Proprioceptors Position of the body in space The sensation of feet
and body parts in relation to on the ground
each other
Note: The receptor neuron cell along with the non-neural tissue is
known as the sensory organ.
B. Based on what they detect :
Examples:
Type Sense Examples/receptors
Receptor Sense Organ Mechano-receptors Mechanical deformation of tissue See : further classification of
Hearing Hair cell Organ of Corti the mechanoreceptors below
Muscle length Nerve endings of Ia Muscle spindle Nociceptors Noxious stimuli i.e. those stimuli that Pain, via free nerve endings
and II afferents cause physical/chemical damage to tissues
Muscle tension Nerve endings of Ib Golgi Tendon Electromagnetic receptors Light waves Rods and cones
afferents Organ or GTO Chemoreceptors Chemical changes Taste, smell
Arterial pO2 Glomus cells* Carotid and aortic Thermoreceptors Temperature changes in the tissues Warm, cold
bodies*
The mechanoreceptors can further be sub-classified as under :
*(You would have learnt about them in the lecture on
regulation of respiration)
 1. Tactile/cutaneous (i.e. based on touch; e.g. touch, pressure, very slowly. In other words, such receptors are designed to sense as
vibration etc.) long as the stimulus is there.
2. Deep tissue sensibility e.g. free nerve endings, Merkel’s disc,
Ruffini’s endings, Pacinian corpuscles
III. Properties of receptors:
3. Muscles, tendons, joints e.g. muscle spindle, Golgi tendon
organ 1. Transducer function (the concept has already been discussed
4. Hair cells for hearing (organ of Corti) above):
5. Hair cells for equilibrium (Vestibular apparatus) The events are best studied in the Pacinian corpuscles (because they
are large, accessible and easy to dissect):
6. Blood pressure:* Pacinian corpuscles:
a) Arterial : baroreceptors in carotid sinus and aortic Site: Subcutaneous tissue, mesentery
arch Structure:
b) Central venous pressure : receptors in the great veins Consists of
and atria i) Unmyelinated nerve
The tactile receptors can in turn be classified as endings/terminal
ii) Surrounded by
i) Rapidly adapting concentric lamellae of
connective tissue
Rapidly adapting Slowly adapting
(capsule)
Also known as Phasic receptors Tonic receptors
Sense Rate of change of stimulus Steady stimulus iii) Nodes of Ranvier : the
Sense Vibration Pressure first node of Ranvier is
Structure These have encapsulated These have expanded endings inside the lamellae and
endings the second node of
Examples Pacinian corpuscles, Meissner’s Merkel’s disc, Ruffini’s Ranvier is just outside
corpuscles, Krause’s end bulb endings, C-mechanoreceptors the lamellae
ii) Slowly adapting:
iii) Adaptationat the level of receptors means:

When a continuous, constant strength stimulus is applied to a

receptor, the response decreases. It is logical to presume that those
receptors that are meant for sub-serving vital functions and
functions necessary for survival (an easy example to recall would be
nociceptors, the receptors for pain) should not adapt or should adapt
 there is receptor
potential and action
potential but no
adaptation

2. Receptor specificity/Adequate stimulus

A receptor is most sensitive (i.e. has the least threshold)
to a particular stimulus; this stimulus is known as its
adequate stimulus. For example, rods and cones are most
sensitive to light (their adequate stimulus) but they also
Study: respond to severe pressure (or blow) on the eyeball; this
Distortion of the membrane  opens sodium channels  causes gives the sensation of flashes of light known as
depolarization  generates receptor or generator potential  if the ‘phosphene’
receptor potential is large enough (which in turn depends on the 3. Adaptation
intensity of the stimulus)  it produces action potential 4. Receptive field of a receptor
The entire area from where the receptor can be
stimulated is known as its receptive field. Larger the
density of the receptors, smaller will be the receptive
Event Site of origin/site Proof
field and better will be the resolution.
responsible
Generator or receptor Unmyelinated nerve Removing the
potential (local endings unmyelinated nerve IV. Laws/Principles of sensory
potential) endings : there is no physiology:
receptor potential and
hence there is no action
potential 1. Muller’s doctrine of specific nerve energies:
Action potential First node of Ranvier Remove/block the first Each nerve carries a specific sensation; stimulating it
node of Ranvier; there anywhere will give this specific sensation.
is receptor potential but Each sensory nerve ends at a particular location in the
no action potential sensory cortex; that particular location sub-serves one
Adaptation Lamellae Remove the lamellae: particular sensation . Therefore, stimulating the nerve
 anywhere along its path produces that particular
sensation only.
2. Law of Projection
Sensation is ‘projected’ by the brain to the site of the
receptor. Therefore, stimulating the nerve anywhere in
its pathway produces the sensation at the site of the
receptor.
Clinical significance:
Law of projection helps in explaining the ‘phantom limb’.

Phantom limb:

Now, when the exposed part of the nerve gets stimulated, the
sensation (say, pain) gets projected to the amputated hand 
therefore, the patient feels pain in the amputated or the missing
hand. In other words, he feels pain in the hand which is not there 
this is known as phantom limb. It is explained on the principle of
law of projection.

1. Encoding of the intensity of the stimulus

Meaning: How is the intensity of the stimulus coded? In
other words, how does one feel the severity of a
sensation e.g. mild pain or severe pain?
If the intensity of the stimulus is more  then,
The intensity of a sensation can be encoded in two ways: more number of receptors will be stimulated 
and the severity of the sensation will be more.
i) By the number of receptors that have been
stimulated:
ii) By the frequency of the action potentials:
2.
If the intensity of the stimulus is more  the amplitude of the
receptor potential will be more  the frequency of the action
potential will be more  and the severity of the sensation will be
more.
This is further clarified by the diagrams below:
a) Example ‘A’ : when the amplitude of the
receptor potential is large; here the
frequency of the action potential is more :
In the diagram, the amplitude of the receptor potential is shown in
blue colour.
The action potential occurs when the magnitude of the receptor
potential reaches the threshold level; it continues to occur
repeatedly as long as the receptor potential is above this threshold.
However, the next action potential can only occur after the
refractory period of the previous action potential i.e. when the
repolarization reaches below the refractory period.
b) Example ‘B’ : when the amplitude of the
receptor potential is small; here the
frequency of the action potential is less:
4. Mathematical laws relating subjective sensation to the intensity
of the stimulus:
i) Weber- Fechner’s law
ii) Stevens’ power law
Introduction:
The intensity of the stimulus (say ‘I’) is the physical correlate
(which can be quantified/measured) whereas the sensation (say
‘S’)is a psychological correlate (which cannot be
quantified/measured). For example, temperature is a physical
correlate, which can be accurately measured (say 21.7 degree
Celsius) whereas the feeling of warmth/cold is the psychological
correlate; the latter is subjective and cannot be quantified. It can
only be expressed as very cold, very warm etc.
Many psychophysicists carried out experiments to find out if they
could derive any mathematical relationship between ‘I’ and ‘S’:
Weber-Fechner law
This relates the sensation felt to the intensity of the stimulus by a
logarithmic function:
S = k log I
Stevens’ power law
As the name suggests, this relates the sensation felt to the intensity
of the stimulus by a power function:
S = k Ia
Where ‘k’ and ‘a’ are constants for a particular stimulus.

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